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Redalyc.Patterns of Leaf Epicuticular Waxes in Species of Clusia Interciencia ISSN: 0378-1844 [email protected] Asociación Interciencia Venezuela Medina, Ernesto; Aguiar, Guillermina; Gómez, Matilde; Medina, José D. Patterns of leaf epicuticular waxes in species of clusia: taxonomical implications Interciencia, vol. 29, núm. 10, octubre, 2004, pp. 579-582 Asociación Interciencia Caracas, Venezuela Available in: http://www.redalyc.org/articulo.oa?id=33909707 How to cite Complete issue Scientific Information System More information about this article Network of Scientific Journals from Latin America, the Caribbean, Spain and Portugal Journal's homepage in redalyc.org Non-profit academic project, developed under the open access initiative PATTERNS OF LEAF EPICUTICULAR WAXES IN SPECIES OF Clusia: TAXONOMICAL IMPLICATIONS Ernesto Medina, Guillermina Aguiar, Matilde Gómez and José D. Medina SUMMARY The genus Clusia L. (Clusiaceae) encompasses ca. 300 species quantity of hexane-soluble compounds extractable from the leaf sur- and occurs from southern USA and Mexico, to southern Brazil and face, which amount to >90% in Clusia rosea, C. orthoneura, and C. Bolivia. It includes free-standing trees and shrubs, hemiepiphytes, minor, the presence of the triterpenes α-amyrin and lupeol in C. epiphytes, and lianas. Taxonomic analysis of this genus is difficult multiflora, and of friedelin and taraxerol, together with C33 and because of the poor preservation of floral material after drying. C35, in C. grandiflora and C. schomburgkiana. The results suggest This work explores the composition of epicuticular waxes in order that the relative proportions of alkanes and triterpenoids in epicu- to allow characterization, at the species level, using chemical mark- ticular waxes may have taxonomic significance for separating spe- ers. The six species analyzed could be separated using the relative cies or infrageneric sections. RESUMEN El género Clusia L. (Clusiaceae) comprende unas 300 Las especies pudieron separarse en base a la proporción de especies que ocurren desde México y el sur de EEUU hasta alcanos, >90% del total en Clusia rosea, C. orthoneura y C. mi- Bolivia y el sur de Brasil. Entre ellas se incluyen árboles y nor, a la presencia de los triterpenos α-amirina y lupeol en C. arbustos, hemiepifitas, epifitas y lianas. El análisis taxonómico multiflora, y de friedelina y taraxerol, conjuntamente con C33 y del género se dificulta por la pobre preservación de las flores al C35 en C. grandiflora y C. schomburgkiana. Los resultados ser secadas. Este trabajo explora la composición de ceras sugieren que la proporción de alcanos y triterpenoides de ceras epicuticulares para caracterizar especies mediante marcadores epicuticulares tiene importancia taxonómica y puede ser utiliza- químicos. Se analizó la composición del extracto obtenido de da para separar especies o secciones infragenéricas. seis especies mediante lavado de la superficie foliar con hexano. Introduction Clusia species possess charac- (Bittrich and Amaral, 1996; etration of liquid water into teristics possibly associated Marsaioli et al., 1999). intercellular spaces, and The genus Clusia includes with tolerance to dry condi- Epicuticular waxes play a avoiding the establishment of around 300 species that occur tions (succulent and/or leathery role in plant biology in water epiphyllic organisms (Nein- throughout the interneotropi- leaves, low leaf conductance; loss regulation, because cu- huis and Barthlott, 1997). cal realm, from southern USA Lüttge, 1996; Pipoly et al., ticular transpiration is related The composition of epicu- and Mexico to southern Brazil 1998). Several species have to the permeability of the cu- ticular waxes is dominated by and Bolivia (Pipoly et al., been shown to be constitutive ticle to water vapor and to long- (>C21), frequently odd- 1998). It includes trees and CAM, or switching from C3 the habitat in which plants chain alkanes (Barthlott, shrubs that grow as free- to CAM metabolism under grow (Schreiber and Riederer, 1989). Other related com- standing individuals on a vari- drought conditions (Franco et 1996). Shady and/or humid pounds are very long chain ety of substrates (shallow clay al., 1990; Lüttge, 1996). habitats frequently produce alcohols and acids, and in or deep sandy soils, calcare- Many Clusia species pro- leaves more permeable to wa- many species triterpenoids ap- ous or serpentinitic soils), or duce resiniferous waxes in the ter vapor than leaves from pear in amounts usually as hemiepiphytes, epiphytes staminate, and/or the pistillate dry and/or sunny exposed small. The amount of epicu- and lianas. flowers, providing material for habitats (Bondada et al., ticular waxes in a given plant All species produce latex honey bees nest construction 1996). In addition, epicuticu- population may be used to varying in abundance, density, (Cuesta-Rubio, 2002). Produc- lar waxes act as insulators identify ecological conditions and color (Engler, 1925; tion of these resins is signifi- from excess environmental (drought and sun exposure), Pipoly et al., 1998). Numerous cant as a pollinator attractant humidity, preventing the pen- while its composition may KEYWORDS / Alkanes / Chemotaxonomy / Clusia / Epicuticular Waxes / Triterpenes / Received: 04/12/2004. Modified: 08/23/2004. Accepted: 08/26/2004. Ernesto Medina. Biologist, Uni- Centro de Ecología. IVIC. nal, Center for Ecology, José D. Medina. Chemist, versidad Central de Venezuela Apartado 21827. Caracas 1020- IVIC, Venezuela. UCV, Venezuela. Doctor in (UCV). Doctor in Agronomy, A. Venezuela. Matilde Gómez. Chemist, Uni- Chemistry, University of University of Hohemheim, e-mail: [email protected] versidad Simón Bolívar, Ven- Laval, Canada. Researcher, Stuttgart Germany. Professor, Guillermina Aguiar. Chemist, ezuela. Associate Research Center for Chemistry, IVIC, UCV and Researcher, Instituto Instituto Pedagógico de Professional, Center for Chem- Venezuela. Venezolano de Investigaciones Caracas, Venezuela. Asso- istry, IVIC, Venezuela. Científicas (IVIC). Address: ciate Research Professio- OCT 2004, VOL. 29 Nº 10 0378-1844/04/10/579-04 $ 3.00/0 579 RESUMO O gênero Clusia L. (Clusiaceae) compreende umas 300 es- em base à proporção de alcanos, >90% do total em Clusia pécies que ocorrem desde México e o sul dos EE.UU. até Bolí- rosea, C. orthoneura e C. minor, à presença dos triterpenos α- via e o sul do Brasil. Entre elas se incluem árvores e arbustos, amirina e lupeol em C. multiflora, e de friedelina e taraxerol, hemiepífitas, epífitas e lianas. A análise taxonômica do gênero conjuntamente com C33 e C35 em C. grandiflora e C. se dificulta pela pobre preservação das flores ao ser secadas. schomburgkiana. Os resultados sugerem que a proporção de Este trabalho explora a composição de ceras epicuticulares para alcanos e triterpenóides de ceras epicuticulares tem importância caracterizar espécies mediante marcadores químicos. Se analisou taxonômica e pode ser utilizada para separar espécies ou seções a composição do extrato obtido de seis espécies mediante lavado infragenéricas. da superfície foliar com hexano. As espécies puderam separar-se help in the identification of Results ecologic and taxonomic groups. The latter is possible Amount of hexane-soluble as the result of the conserva- compounds extracted tive biochemical pathways that lead to wax synthesis The average load of hex- (Bianchi, 1987; Gülz, 1994). ane-soluble compounds (HSC; Extraction and identification Figure 1 for adult leaves) was of wax compounds is rela- relatively higher in C. schom- tively straightforward using burgkiana, C. grandiflora, and volatile low polarity organic C. orthoneura (≥20µg·cm-2), solvents, gas chromatography than in C. multiflora and C. and mass spectrometry. This minor (<20µg·cm-2). On the paper reports on epicuticular abaxial side only C. ortho- wax profiles of five Clusia neura and C. schomburgkiana species in order to test the had values ≥20µg·cm-2. The feasability to a) separate spe- -2 amount of HSC on the cies using the relative propor- Figure 1. Total hexane-soluble compounds (µg·cm ) extracted from the adaxial leaf side of adult tions of alkanes and presence/ surfaces of adult leaves of Clusia species. Under the Abaxial and Adaxial leaves was larger than on the headings bars followed by the same letter are not statistically different. absence of triterpenoids; b) abaxial side in C. rosea and evaluate variations of wax C. grandiflora, while the op- load and composition in rela- Epicuticular waxes were 4ºC/min up to 320ºC and posite occurred in C. schom- tion to leaf age and leaf side, extracted from at least three maintained for 10min. The in- burgkiana and C. orthoneura. and the influence of ecologi- fresh leaf replicates, separat- jector was maintained at In C. multiflora and C. minor cal conditions such as sun ex- ing adaxial and abaxial sides 250ºC and the detector at differences in HSC content on posure. of young and mature, healthy 320ºC. Results are reported as both sides were small. In leaves. Leaf area was mea- amount of extracted waxes young leaves these relation- Material and methods sured before extraction (Licor per unit leaf area, and as rela- ships were similar. No statisti- LI-3100 area meter), and leaf tive proportions as indicated cal differences could be es- Clusia multiflora HBK dry weight was determined by the average percentage of tablished between leaves of (Section Anandrogyne) and after extraction (ventilated chromatogram area for each different ages, or between leaf C. minor L. (Section Retin- oven at 60ºC until constant chemical species. Compound
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